Somatic growth and otolith microstructure of larval and juvenile pacific cod <Emphasis Type="Italic">Gadus macrocephalus</Emphasis>

ABSTRACT:   Microstructures of sagittae and lapilli were examined in relation to somatic growth for reared larvae and juveniles of Pacific cod. The Laird–Gompertz model was fitted to the daily age and somatic growth relationship. Growth increments were deposited on a daily basis in both kinds of otoliths, with a check formed at hatching. Two subsequent checks and an accessory primordia (AP) occurred in the sagittae. The lapillus was adequate for increment width measurement through the early life stages. Sagittal and frontal plane of sagitta was adequate for measurement in the pre-AP and post-AP formation stages, respectively. The shift of desirable plane was caused by changes in otolith and increment shapes with AP formation. Back-calculated total lengths using the biological intercept method did not significantly differ with certain body lengths ( P  > 0.05), suggesting validity of back-calculation in this species. Using the back-calculated total length, morphological and ecologic changes that seemed to affect checks and AP formations are discussed.     

Otolith development and daily increment formation in laboratory-reared larval and juvenile black-spot tuskfish <Emphasis Type="Italic">Choerodon schoenleinii</Emphasis>

Microstructures of lapilli were examined for reared larvae and juveniles of black-spot tuskfish Choerodon schoenleinii. Lapilli of larvae at 1 day after hatching have one diffuse and obscure ring with an otolith radius of 4.3 ± 0.50 μm (mean ± SD, N = 8). The slope and intercept of the regression between the number of days after hatching and increment counts did not differ significantly from one and zero, respectively, indicating that lapillus increments were formed on a daily basis after hatching. There was an ontogenetic shift in the relative values of somatic and otolith growth, which corresponded to the transition from pelagic larvae to settlement stage. Simultaneously, the daily increment width reached the maximum value. These findings suggest that age at maximum value of increment width can be used as an indicator of the planktonic larval duration while settlement mark is not found. Since ontogenetic shift in the relationship between otolith radius and body size was observed, back-calculation of somatic growth in black-spot tuskfish using the otolith radius during the early life stages should be analyzed with caution, and the method requires further validation.     

叉尾斗鱼仔鱼耳石形态发育与日轮形成特征

对实验室人工繁殖的叉尾斗鱼(Macropodus opercularis)仔鱼耳石形态发育与日轮进行了观察研究.结果显示,叉尾斗鱼微耳石和矢耳石在胚胎时已出现.微耳石在仔鱼刚孵化时为圆盘状,随仔鱼发育转变为近椭圆形,孵出后19 d转变为中部圆凸两端较尖的菱形;矢耳石在仔鱼刚孵出时为圆盘状,随仔鱼发育转变为椭圆形,孵出后19 d转变为一端略尖的桃形;星耳石在仔鱼孵出后第19天才出现,呈中部略凹的椭圆形.叉尾斗鱼仔鱼耳石长径(包括微耳石和矢耳石)与鱼体全长(TL,mm)呈线性相关.仔鱼耳石上第一个轮纹在孵出后第2天形成,其后每日形成1个新轮纹,生长轮数目与仔鱼日龄(D)呈线性相关,且线性方程斜率接近于1.结论认为,叉尾斗鱼仔鱼星耳石出现时间晚,矢耳石形态在后期出现较大变化,而微耳石在胚胎时即形成,形态稳定;日轮可读性较好,故更适合做日轮研究的材料.本研究旨在为叉尾斗鱼自然种群年龄结构调查及其资源保护提供基础数据.     

Early life history of the black anglerfish Lophius budegassa Spinola, 1807 in the Mediterranean Sea using otolith microstructure

The early life history of the black anglerfish, Lophius budegassa was investigated by otolith (lapilli) increment analysis. Samples of demersal juvenile L. budegassa ranging from 54 to 196 mm total length were collected during bottom trawl surveys in the central Adriatic Sea. By counting increments presumed to be deposited daily in the lapillar otoliths, 88 specimens of L. budegassa were successfully aged. Age estimates of juveniles ranged between 79 and 204 days, indicating that probably the pelagic phase of this species is relatively short and settlement occurs at less than 3 months of life. The analysis of check marks in the core area of lapilli enabled us to determine the period of endogenous feeding, which would last between 15 and 24 days after hatching. Back-calculated hatching dates and, consequently, the spawning season of L. budegassa in the Adriatic Sea was spread over a long period, lasting at least from February to June. The length at age relationship gave an estimate of mean growth rate of approximately 0.8–1.02 mm/day, indicating a faster growth rate of 0+ juveniles L. budegassa than previously thought. The implications of these findings on age estimates discrepancies between previous ageing studies on L. budegassa carried out using different calcified structure (sagittae or illicia) are discussed.     

Otolith microstructure of chum salmon <Emphasis Type="Italic">Oncorhynchus keta</Emphasis>: formation of sea entry check and daily deposition of otolith increments in seawater conditions

To apply otolith microstructure to examination of age and growth of juvenile chum salmon Oncorhynchus keta inhabiting coastal waters, formation of otolith increments was investigated for juveniles reared in a seawater aquarium and in net pens. In all otoliths examined, a distinctive check was formed at the time of sea entry of the fish. The deposition of otolith increments after the check was daily for rearing both in the aquarium (57 days) and in the net pens (26 days). Check formation associated with sea entry was also observed in otoliths of juvenile salmon collected 1 km off the coast of Shari, Hokkaido, Japan. Transmitted light observation of otoliths of those fish revealed a transition in otolith increment appearance from dark to light. Otolith Sr: Ca ratio remarkably changed from a low to a high level, coinciding with the transition in otolith appearance. It is suggested that the transition was associated with individual sea entry. This study demonstrated that the check and/or transition associated with sea entry are applicable to a benchmark for otolith increment counts of juvenile chum salmon inhabiting coastal waters.     

Sagittal otolith microstructure,early growth and development of Coilia ectenes in the Yangtze Estuary,China

Larvae and juveniles of Coilia ectenes were collected in the Yangtze Estuary during June through September 2009 and in May 2011. Our objectives were to reveal the otolith microstructural pattern, and then to reveal the early growth and development process through otolith analysis for C. ectenes. A total of 368 individuals, ranging 4.4–81.2 mm body length (BL) and spanning a continuum from yolk-sac larvae to early juveniles, were analyzed. There is a single primordium in the core of the sagittal otolith, surrounded by a prominent dark band (PD-band) and a wide light area (WL-area). Regular bipartite increments deposit outside of the WL-area. Similar to other Engraulidae species, the PD-band forms corresponding to hatching, increments form daily, and the first increment forms on the 3rd day post hatching, corresponding to the first feeding. The relationship of otolith radius and BL was significantly fitted to a piecewise regression with the inflection point at 34.0 mm BL. Mid-age and mid-size of transition from yolk-sac to preflexion was calculated as 5 days and 6.1 mm BL, from preflexion to flexion was 18 days and 15.2 mm BL, and from flexion to postflexion 29 days and 21.5 mm BL. Mid-size from postflexion to juvenile was 26.6 mm BL. These results can provide an essential basis for future studies on early-life ecology of this species.     

大黄鱼仔稚鱼不同发育阶段矢耳石形态发育和微结构特征

对人工培育大黄鱼(Larimichthys crocea)的生长发育与矢耳石形态及微结构特征进行研究,结果表明:(1)大黄鱼矢耳石上的轮纹是每日形成的,第1日轮在孵化后第2天形成,与其初次摄食相对应。(2)大黄鱼卵黄囊期和前弯曲期仔鱼的耳石形态为圆形,进入弯曲期耳石长轴迅速伸长,在后弯曲期耳石形态变为椭圆形。进入稚鱼期,矢耳石开始形成次生核。随后次生核数量逐渐增加,在孵化后47～78 d的个体中,次生核数量稳定在5～7个,耳石近似盾形。(3)根据耳石日轮宽度推算的大黄鱼稚鱼在其仔鱼期生长率(b)与第1个次生核的形成时间(tsp1)之间存在明显的线性关系,表明生长较快的个体形成次生核的时间较早,进入稚鱼期所需的时间更短。以上结论表明,大黄鱼矢耳石可以反演其早期生活史阶段的生长发育特征。     

Faster growth before metamorphosis leads to a higher risk of pseudoalbinism in juveniles of the starry flounder <Emphasis Type="Italic">Platichthys stellatus</Emphasis>, as suggested by otolith back-calculation

The mechanism underlying malformation during flatfish metamorphosis is not clear. Here, the relationship between growth before metamorphosis and the occurrence of malformation after metamorphosis was examined in individuals reared in a single tank. Otoliths of starry flounder Platichthys stellatus larvae were labeled with alizarin complexone before metamorphosis [12 days after hatching (DAH)]. The positive linear regression between the lapillus otolith diameter and body length at 12 DAH enabled the estimation of individual larval size after completion of metamorphosis. At 12 DAH, the average body length of the typical pseudoalbino juveniles was significantly greater (5.37 mm) than that of the normal juveniles (5.13 mm) (P < 0.01, n = 100). In addition, of the larvae that were estimated to measure less than 4.8 mm at 12 DAH, approximately 80% metamorphosed into normal juveniles, whereas more than 60% of the larvae that measured more than 5.4 mm became pseudoalbino juveniles. These results indicate that the larvae that grow rapidly during the first 12 days are more likely to become pseudoalbino individuals after metamorphosis.     

Growth of juvenile chub mackerel Scomber japonicus in the western North Pacific Ocean: with application and validation of otolith daily increment formation

The growth of juvenile chub mackerel Scomber japonicus collected in the western North Pacific Ocean in 2007 and 2009 was examined based on the evidence of otolith daily increment formation in captive specimens. There was a significant difference in the relationship between known age and number of increments in the frontal and sagittal planes. Repeated markings on the otolith using Alizarin complexone and the coefficient of variation in number of increments suggest that the increments in the frontal plane of the otolith are more suitable for age estimation than those in the long and short axes of the sagittal plane. The increments in the frontal plane formed daily, and the first ring was usually deposited 3 days after hatch. Age of wild juveniles ranged from 24 to 211 days after hatch based on the frontal plane method. The estimated hatching periods of specimens ranged from February to June, but the April-hatched specimens were collected throughout the sampling periods of 2007 and 2009. The Gompertz growth model showed a difference in growth pattern in specimens between 2007 and 2009. The juveniles in 2009 appeared to grow more quickly than those in 2007 until summer, but thereafter the 2009 specimens seemed to grow more slowly.     

月鳢仔鱼耳石的荧光标记及其日轮确证

利用茜素络合物对体长(9.99±0.56)mm的月鳢(Channa asiatica)仔鱼耳石进行浸泡标记试验。结果表明,经荧光显微镜检测,80 mg/L、100 mg/L和120 mg/L的溶液浸泡24 h后均可以在月鳢仔鱼的耳石上形成橘红色标记环,星耳石、微耳石和矢耳石的标记率均为100%;其中,以星耳石标记环荧光强度最大,且在普通光学显微镜下也可观察到紫红色标记环。经100 mg/L和120 mg/L溶液浸泡后,月鳢仔鱼在其后饲养阶段出现较高的死亡率,显示高浓度茜素络合物溶液对月鳢仔鱼有一定毒性。标记后星耳石上的新增轮纹数(N)和饲养天数(D)密切相关,其线性方程为:N=0.9291D-0.2974(R2=0.9793,n=64,P<0.01),斜率0.9291与1无显著差异(P>0.05),表明星耳石上的生长轮即为日轮。研究表明,在追踪月鳢仔鱼迁移行为时,可采用80 mg/L的茜素络合物溶液对其星耳石进行染色标记。     

The effect of docosahexaenoic and eicosapentaenoic acids in live food on the development of abnormal morphology in hatchery-reared brown sole Pseudopleuronectes herzensteini

We examined the effect of docosahexaenoic acid (DHA) and eicosapentaenoic acid (EPA) on the rates of abnormal morphology in juvenile brown sole Pseudopleuronectes herzensteini. Larvae during the D–E stages (15–24 days post hatching) were fed live food containing various amounts of DHA and/or EPA prepared using emulsified oils (DHA ethyl ester, EPA ethyl ester, and corn oil). Larvae during the F–I stages were fed Artemia enriched with a commercial diet supplement. We found that DHA and EPA promoted larval development and improved the incidence of morphological abnormalities in brown sole juveniles to a similar extent. However, DHA was more effective than EPA in preventing the appearance of morphological abnormalities in brown sole. The incidence of normal morphology was clearly improved by an increase of the DHA content in brown sole larvae at 25 days post-hatching. These results suggest that it is important to promote larval development and feed larvae with live food containing high levels of DHA during the D–E stages to prevent morphological abnormalities in brown sole juveniles.     

Lack of essential fatty acids in live feed during larval and post-larval rearing: effect on the performance of juvenile <Emphasis Type="Italic">Solea senegalensis</Emphasis>

Despite the large progress obtained in recent years, Senegalese sole (Solea senegalensis) production of high quality juveniles is still a bottleneck. This paper examines the effect of larval and post-larval lipid nutrition on juvenile performance and quality. Four dietary treatments were tested: A—enriched Artemia spp. (EA); B—non-enriched Artemia spp. (NEA); C—EA during the pelagic larval period and NEA after larval settlement; D—50% EA and 50% NEA. Juvenile fatty acid profile at 60 days after hatching (DAH) clearly reflected the larval and post-larval diet composition. Feeding sole larvae on NEA (poor in lipids and essential fatty acids-EFA) had a negative effect, reducing growth (total length and dry weight) after 30 DAH and decreasing digestive enzyme activity at the end of the rearing period (60 DAH). However, relatively good performance compared to the EFA-richest treatment (A) was obtained when larvae were fed 50% EA and 50% NEA (D) or even EA only during the pelagic larval period followed by NEA after larval settlement (C). Malpigmentation was not affected by the dietary regimes and its incidence was very low. However, skeletal deformities were prevalent, particularly in the caudal complex, independently of diet. The results confirm that Senegalese sole appear to have lower larval EFA requirements than most cultured marine species and potentially even lower requirements during the post-larval stage. The importance of studying the impact of early nutrition on later juvenile stages was clearly highlighted in this study.     

Daily and sub-daily otolith increments of larval and juvenile walleye pollock, Theragra chalcogramma (Pallas), as validated by alizarin complexone experiments

Walleye pollock (Theragra chalcogramma) were reared from eggs to the juvenile life stage to study daily increment formation in the sagittae otoliths, which are routinely used for age and growth analyses. The apparent deposition of sub-daily growth increments becomes problematic for determining fish age from the late larval stage throughout the juvenile (young-of-the-year) development stage. Otolith marking experiments were conducted to determine interpretation criteria to differentiate between daily and sub-daily increments. Immersion of larval and transforming walleye pollock in 25 mg/l of alizarin complexone (ALC) for 6 h once a week produced a fluorescent mark on the day of staining. Evidence of six well defined and equally spaced increments counted between the weekly ALC marks validated the deposition of daily increments. The daily increments gradually increased in width as the fish/otolith grew. The criteria for determining the presence of sub-daily increments between the daily increments were (1) weak optical definition and (2) a sudden change in incremental distance that lasted for one or two increments and were approximately <0.5 μm in width. Growth problems that occurred during the experiments were identified on otoliths as reductions in daily incremental widths and optical definition, which continued for several days. Otoliths from field-collected fish have also shown similar changes in daily increment properties during the juvenile stage, which may be an indicator of an environmental influence. The criteria for defining different increment types help to resolve our current age determination issues for late larval and early juvenile stage walleye pollock from the Gulf of Alaska.     

Larval and juvenile Australian smelt Retropinna semoni somatic and otolith growth parameters – implications for growth interpretation of wild fish

Abstract –  Relationships between fish length, otolith size, age and weight were assessed for a population of wild Australian smelt ( Retropinna semoni ) larvae and juveniles captured over a 4-year period to aid further interpretation of growth and condition during the early life history of the species. Nonlinear smoothed generalized additive models best described the fish–otolith size relationship during the larval and juvenile period, indicating that the proportionality between fish length and otolith size varies in relation to size. It is proposed that back-calculated predictions of fish size at a previous age or otolith size, accounting for individual variation is possible by assuming a body proportional hypothesis. Growth rate of larval and juvenile Australian smelt was best described using the Gompertz model that indicated a steady decline in growth rate after around 30 days of age. The allometric growth of larval and juvenile Australian smelt established from the length/weight relationship can subsequently be used to assess the condition of fish within this population using a relative condition or relative weight condition index. The results of the study have provided significant information to enable more precise growth reconstruction and condition assessment for the species in Australian lowland rivers.     

Feeding rhythm and growth of the tongue sole, Cynoglossus semilaevis Günther, during its early life stages

The feeding rhythm and growth characteristics of the early life stages of the tongue sole, Cynoglossus semilaevis Günther, were studied. Larvae began to prey on rotifers about 2–3 days after hatching at 23°C. About 12 days after hatching, larvae had grown to 8–9 mm in total length and were able to prey on Artemia larvae. As the larvae grew, they showed an increasing feeding capacity and a distinct feeding rhythm. Feeding intensity for day‐6 larvae was highest at 12:00 and 18:00 hours, about 2–3 h after the maximum feeding incidence. The highest levels of feeding intensity for day‐16 larvae occurred between 09:00 and 18:00 hours. By day 26, when the larvae had metamorphosed, feeding capacity had again increased considerably and, in contrast to the earlier stages, feeding intensity peaked at 18:00 and 24:00 hours concurrently with feeding incidence. Thus, tongue sole were found to have different feeding rhythms in the pre‐ and post‐metamorphosis stages, with the highest feeding activity in the daytime during the larval planktonic stage, and at night during the juvenile benthic stage. The estimated maximum daily feeding rates were 65%, 40% and 11% of body weight on days 6, 16 (larvae) and 26 (juveniles) respectively. Size variation increased markedly with development.     

茜素络合物对鲤仔鱼耳石标记特征研究

利用100 mg/L的茜素络合物（alizarin complexone,ALC）对鲤仔鱼进行48 h的水环境浸泡标记,以探讨该ALC标记方法的特征,及其对矢耳石、星耳石和微耳石的标记效果以及鱼体ALC浸泡、续养恢复与耳石ALC标记区域形成和消失的时滞进行研究。结果显示,3种耳石在可见光和荧光下均能检测到明显的标记环。其中星耳石的标记效果最佳,微耳石次之。耳石上荧光信号出现和消失的时间与鱼体ALC浸泡开始和结束的时间均存在1 d的时滞。此外,浸泡标记过的鲤仔鱼在进行了长达50 d的续养恢复后,其耳石上的ALC标记环仍清晰可见。研究表明,ALC标记法所形成的标记环在耳石上可长期存在,使用ALC对鲤仔鱼进行生态标记具有很强的可行性。     

Rearing dover sole larvae onTisbe andArtemia diets

The suitability of the harpacticoid copepodTisbe holothuriae as a diet for larval and juvenile Dover sole (Solea solea) was assessed by rearing groups of sole for 42 days under a range of dietary regimes. Larval sole, approximately 1 week old, were reared onTisbe, Artemia, or a mixedTisbe-Artemia diet for 13 days. No significant differences in length between sole larvae from any diet were found after this time, but larvae offeredArtemia alone showed a significantly higher frequency of malpigmentation than those offered the other diets. After metamorphosis (day 16), survivors of this experiment were reared for a further 29 days on various diets to give the following dietary sequences:Tisbe-fed larvae, fed onArtemia as juveniles (Tis.-Art.);Artemia-fed larvae, fed onArtemia as juveniles (Art.-Art.);Artemia-fed larvae, fed onTisbe as juveniles (Art.-Tis.) and mixed diet fed larvae, fed on a mixed diet as juveniles (AT-AT). At the end of this period AT-AT and Tis.-Art.-fed juveniles were significantly larger than those on the Art.-Art. dietary regime. Juveniles from the Tis.-Art. dietary regime consumed more prey items than the Art.-Art group. AT-AT juveniles consumed similar amounts of food to Art.-Art. juveniles but were significantly larger after 29 days culture. This was attributed to the presence ofTisbe in their diet. Overall, the best larval and juvenile diet appeared to be a mixed diet throughout the culture period.     

Interannual variability in hatching period and early growth of juvenile walleye pollock, Theragra chalcogramma, in the Pacific coastal area of Hokkaido

Juvenile walleye pollock of the Japanese Pacific population were collected from the Funka Bay [spawning ground; 16–64 mm fork length (FL)] in spring and the Doto area (nursery ground; 70–146 mm FL) in summer. Hatch dates were estimated by subtracting the number of otolith daily increments from sampling dates, and their early growth was back‐calculated using otolith radius–somatic length relationships. Interannual change of the hatching period was observed during 2000–02, and the peaks ranged from mid‐February in 2000 to early‐April in 2002. In 2000, when a strong year class occurred, early life history of the surviving juveniles could be characterized by early hatching and slower growth in the larval stage (<22 mm length). Higher growth rate in 2001 and 2002 did not always lead to good survival and recruitment success. Even though their growth was slow in 2000, the larvae hatched early in the season had larger body size on a given date than faster‐growing larvae hatched in later season in 2001 and 2002. Bigger individuals at a certain moment may have advantage for survival. The delay of hatching period may result in higher size‐selective mortality, and as a necessary consequence, back‐calculated growth in 2001 and 2002 could shift towards higher growth rate, although abundance of such a year class would be at the lower level. Variability in spawning period, early growth and their interaction might have a strong relation to larval survival through cumulative predation pressure or ontogenetic changes in food availability.     

Interannual differences in growth and hatch‐date distributions of early juvenile European anchovy in the Bay of Biscay: implications for recruitment

In order to understand better the recruitment variability in European anchovy in the Bay of Biscay, it is important to investigate the processes that affect survival during the early life stages. Anchovy juvenile growth trajectories and hatch‐date distributions were inferred over a 3‐year period based on otolith microstructure analysis. Otolith growth trajectories showed a characteristic shape depending on their hatch‐date timing. Earlier‐born juveniles had notably broader maximum increments than later born conspecifics, resulting in higher growth rates. This observation suggests that early hatching would be beneficial for larval and juvenile growth, and, therefore, survival. The estimated juvenile hatch‐date distributions were relatively narrow compared with the extended anchovy spawning season (March–August) in the Bay of Biscay and indicated that only individuals originated mainly from the summer months (June–August) survived until autumn. Hatch‐date distributions were markedly different among years and seemed to influence the interannual recruitment strength of anchovy. We conclude that years characterized by juvenile survivors originating from the peak spawning period (May and June) would lead to considerable recruitment success. Downwelling events during the peak spawning period seem to affect larval survival. Furthermore, size‐dependent overwinter mortality would be an additional process that regulates recruitment strength in the anchovy population in the Bay of Biscay.     

云纹石斑鱼胚胎发育及仔、稚、幼鱼形态观察

对云纹石斑鱼Epinephelus moara胚胎发育及仔、稚、幼鱼形态进行了观察与研究,详细描述了各发育期的形态特征和发育时间。结果表明:1)在水温22±0.2℃、盐度30、溶氧7.8mg/L、pH8.25的条件下,云纹石斑鱼的受精卵历时38h 17min开始孵化出膜。胚胎发育可分为受精卵、卵裂、原肠、神经胚和器官形成及出膜6个阶段,受精、胚盘形成、2细胞等28个时期。2)在水温23±1℃,盐度30±3,DO≥5mg/L,pH8.0±0.5的海水中,培育至5d,卵黄囊完全消失,成为后期仔鱼;培育至27d,发育最快的云纹石斑鱼结束仔鱼期,进入稚鱼期;培育至65d,发育最快的稚鱼完成变态,成为幼鱼。胚后发育过程主要是根据卵黄囊、鳍膜、鳞片、体色及第1腹鳍棘与第2背鳍棘相对长度的变化分为仔鱼、稚鱼和幼鱼3个时期。其中仔鱼期又根据其卵黄囊的有无划分为前期仔鱼和后期仔鱼。